plant science research

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Lettuce Gone Wild, part two

The lettuce we eat is a close relative to the lettuce we weed out of our gardens. Last week we discussed the potential that wild relatives may have for improving cultivated lettuce. But if wild lettuce can be crossed with cultivated lettuce to create new cultivars, can cultivated lettuce cross with wild lettuce to make it more weedy?

Because so many of our crops are closely related to some of the weeds found along with them or the plants growing in nearby natural areas, the creation of crop-wild hybrids has long been a concern. This concern is heightened in the age of transgenic crops (also known as GMOs), for fear that hybrids between weeds and such crops could create super weeds – fast spreading or highly adapted weeds resistant to traditional control methods such as certain herbicides. To reduce this risk, extensive research is necessary before such crops are released for commercial use.

flowers of prickly lettuce (Lactuca serriola)

There are no commercially available, genetically modified varieties of cultivated lettuce, so this is not a concern when it comes to crop-wild hybrids; however, due to how prevalent weedy species like prickly lettuce (Lactuca serriola) are, hybridization with cultivated lettuce is still a concern. So, it is important to understand what the consequences might be when hybridization occurs.

In a paper published in Journal of Applied Ecology in 2005, Hooftman et al. examined a group of second-generation hybrids (L. sativa x L. serriola), and found that the hybrids behaved and appeared very similarly to non-hybrid prickly lettuce. They also found that the seeds produced by the hybrids had a significantly higher germination rate than non-hybrid plants. This is an example of hybrid vigor. Thus, “if hybridization does occur, this could lead to better performing and thus potentially more invasive (hybrid) genotypes.” However, the authors cautioned that “better performing genotypes do not automatically result in higher invasiveness,” and that much depends on the conditions they are found in, the level of human disturbance, etc.

Another thing to consider is that hybrids are not stable. In an article published in Nature Reviews Genetics in 2003, Stewart et al. adress the “misunderstanding that can arise through the confusion of hybridization and … introgression.” It is wrong to assume that hybrids between crops and wild relatives will automatically lead to super weeds. For this to occur, repeated crosses with parental lines (also known as backcrossing) must occur, and “backcross generations to the wild relative must progress to the point at which the transgene [or other gene(s) in question] is incorporated into the genome of the wild relative.” That is what is meant by “introgression.” This may happen quickly or over many generations or it may never happen at all. Each case is different.

prickly leaf of prickly lettuce (Lactuca serriola)

In a paper published in Journal of Applied Ecology in 2007, Hooftman et al. observe the breakdown of crop-wild lettuce hybrids. They note that “fitness surplus through [hybrid vigor] will often be reduced over few generations,” which is what was seen in the hybrids they observed. One possible reason why this occurs is that lettuce is predominantly a self-crossing species; outcrossing is rare, occurring 1 – 5% of the time thanks to pollinating insects. But that doesn’t mean that a stable, aggressive genotype could never develop. Again, much depends on environmental conditions, as well as rates of outcrossing and other factors relating to population dynamics.

A significant expansion of prickly lettuce across parts of Europe led some to hypothesize that crop-wild hybrids were partly to blame. In a paper published in Molecular Ecology in 2012 Uwimana et al. ran population genetic analyses on extensive data sets to determine the role that hybridization had in the expansion. They concluded that, at a level of only 7% in wild habitats, crop-wild hybrids were not having a significant impact. They observed greater fitness in the hybrids, as has been observed in other studies (including the one above), but they acknowledged the instability of hybrids, especially in self-pollinating annuals like lettuce.

seed head of prickly lettuce (Lactuca serriola)

It is more likely that the expansion of prickly lettuce in Europe is due to “the expansion of favorable habitat as a result of climate warming and anthropogenic habitat disturbance and to seed dispersal because of transportation of goods.” Uwimana et al. did warn, however, that “the occurrence of 7% crop-wild hybrids among natural L. serriola populations is relatively high [for a predominantly self-pollinating species] and reveals a potential [for] transgene movement from crop to wild relatives [in] self-pollinating crops.”

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Lettuce Gone Wild, part one

Lettuce, domesticated about six thousand years ago in a region referred to as the Fertile Crescent, bears little resemblance to its wild ancestors. Hundreds of years of cultivation and artificial selection eliminated spines from the leaves, reduced the latex content and bitter flavor, shortened stem internodes for a more compact, leafy plant, and increased seed size, among several other things. The resulting plant even has a different name, Lactuca sativa (in Latin, sativa means cultivated). However, cultivated lettuce remains closely related to its progenitors, with whom it can cross to produce wild-domestic hybrids. For this reason, there is great interest in the wild relatives of lettuce and the beneficial traits they offer.

image credit: wikimedia commons

Crop wild relatives are a hot topic these days. That’s because feeding a growing population in an increasingly globalized world with the threat of climate change looming requires creative strategies. Utilizing wild relatives of crops in breeding programs is a potential way to improve yields and address issues like pests and diseases, drought, and climate change. While this isn’t necessarily a new strategy, it is increasingly important as the loss of biodiversity around the globe threatens many crop wild relatives. Securing them now is imperative.

There are about 100 species in the genus Lactuca. Most of them are found in Asia and Africa, with the greatest diversity distributed across Southwest Asia and the Mediterranean Basin. The genus consists of annual, biennial, and perennial species, a few of which are shrubs or vines. Prickly lettuce (L. serriola), willowleaf lettuce (L. saligna), and bitter lettuce (L. virosa) are weedy species with a wide distribution outside of their native range. Prickly lettuce is particularly common in North America, occurring in the diverse habitats of urban areas, natural areas, and agricultural fields. It is also the species considered to be the main ancestor of today’s cultivated lettuce.

In a paper published in European Journal of Plant Pathology in 2014. Lebeda et al. discuss using wild relatives in lettuce breeding and list some of the known cultivars derived from crosses with wild species. They write that in the last thirty years, “significant progress has been made in germplasm enhancement and the introduction of novel traits in cultivated lettuce.” Traditionally, Lactuca serriola has been the primary source for novel traits, but breeders are increasingly looking to other species of wild lettuce.

bitter lettuce (Lactuca virosa) – image credit: wikimedia commons

Resistance to disease is one of the main aims of lettuce breeders. Resistance genes can be found among populations of cultivated lettuce, but as “extensive screening” for such genes leads to “diminishing returns in terms of new resistance,” breeders look to wild lettuce species as “sources of new beneficial alleles.” The problem is that there are large gaps in our knowledge when it comes to wild lettuce species and their interactions with pests and pathogens. Finding the genes we are looking for will require “screening large collections of well defined wild Lactuca germplasm.” But first we must develop such collections.

In a separate paper (published in Euphytica in 2009), Lebeda et al. discuss just how large the gaps in our understanding of the genus Lactuca are. Beginning with our present collections they found “serious taxonomic discrepancies” as well as significant redundancy and unnecessary duplicates in and among gene banks. They also pointed out that “over 90% of wild collections are represented by only three species” [the three weedy species named above], and they urged gene banks to “rapidly [acquire] lettuce progenitors and wild relatives from the probable center of origin of lettuce and from those areas with the highest genetic diversity of Lactuca species” as their potential for improving cultivated lettuce is too important to neglect.

Lactuca is a highly variable genus; species can differ substantially in their growth and phenology from individual to individual. Lebeda et al. write, “developmental stages of plants, as influenced through selective processes under the eco-geographic conditions where they evolved, can persist when plants are cultivated under common environmental conditions and may be fixed genetically.” For this reason it is important to collect numerous individuals of each species from across their entire range in order to obtain the broadest possible suite of traits to select from.

One such trait is root development and the related ability to access water and nutrients and tolerate drought. Through selection, cultivated lettuce has become a very shallow-rooted plant, reliant on regular irrigation and fertilizer applications. In an issue of Theoretical and Applied Genetics published in 2000, Johnson et al. demonstrate the potential that Lactuca serriola, with its deep taproot and ability to tolerate drought, has for developing lettuce cultivars that are more drought tolerant and more efficient at using soil nutrients.

willowleaf lettuce (Lactuca saligna) – image credit: wikimedia commons

Clearly we have long way to go in developing improved lettuce cultivars using wild relatives, but the potential is there. As Lebeda et al. write in the European Journal of Plant Pathology, “Lettuce is one of the main horticultural crops where a strategy of wild related germplasm exploitation and utilization in breeding programs is most commonly used with very high practical impact.”

Coming Up in Part Two: Can cultivated lettuce cross with wild lettuce to create super weeds?

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Highlights from the Alaska Invasive Species Workshop

This October 24-26th I was in Anchorage, Alaska for the 18th annual Alaska Invasive Species Workshop. The workshop is organized by the Committee for Noxious and Invasive Pests Management and University of Alaska Fairbanks Cooperative Extension. It is a chance for people involved in invasive species management in Alaska – or just interested in the topic – to learn about the latest science, policies, and management efforts within the state and beyond. I am not an Alaska resident – nor had I ever been there until this trip – but my sister lives there, and I was planning a trip to visit her and her family, so why not stop in to see what’s happening with invasive species while I’m at it?

What follows are a few highlights from each of the three days.

Day One

The theme of the workshop was “The Legacy of Biological Invasions.” Ecosystems are shaped by biotic and abiotic events that occurred in the past, both recent and distant. This is their legacy. Events that take place in the present can alter ecosystem legacies. Invasive species, as one speaker said in the introduction, can “break the legacy locks of an ecosystem,” changing population dynamics of native species and altering ecosystem functions for the foreseeable future. Alaska is one of the few places on earth that is relatively pristine, with comparably little human disturbance and few introduced species. Since they are at an early stage in the invasion curve for most things, Alaska is in a unique position to eradicate or contain many invasive species and prevent future introductions. Coming together to address invasive species issues and protect ecosystem legacies will be part of the human legacy in Alaska.

The keynote address was delivered by Jamie Reaser, Executive Director of the National Invasive Species Council and author of several books. She spoke about the Arctic and its vulnerability to invasive species due to increased human activity, climate change, and scant research. To address this and other issues in the Arctic, the Arctic Council put together the Arctic Biodiversity Assessment, and out of that came the Arctic Invasive Alien Species Strategy and Action Plan. Reaser shared some thoughts about how government agencies and conservation groups can come together to share information and how they can work with commercial industries to address the threat of invasive species. She stressed that Alaska can and should play a leadership role in these efforts.

Later, Reaser gave a presentation about the National Invasive Species Council, including its formation and some of the work that it is currently doing. She emphasized that invasive species are a “people issue” – in that the actions and decisions we make both create the problem and address the problem – and by working together “we can do this.”

Day Two

Most of the morning was spent discussing Elodea, Alaska’s first invasive, submerged, freshwater, aquatic plant. While it has likely been in the state for a while, it was only recognized as a problem within the last decade. It is a popular aquarium plant that has been carelessly dumped into lakes and streams. It grows quickly and tolerates very cold temperatures, photosynthesizing under ice and snow. It propagates vegetatively and is spread to new sites by attaching itself to boats and float planes. Its dense growth can crowd out native vegetation and threaten fish habitat, as well as make navigating by boat difficult and landing float planes dangerous. Detailed reports were given about how organizations across the state have been monitoring and managing Elodea populations, including updates on how treatments have worked so far and what is being planned for the future. A bioeconomic risk analysis conducted by Tobias Schwörer was a featured topic of discussion.

After lunch I took a short break from the conference to walk around downtown Anchorage, so I missed a series of talks about environmental DNA. I returned in time to hear an interesting talk about bird vetch (Vicia cracca). Introduced to Alaska as a forage crop, bird vetch has become a problematic weed on farms, orchards, and gardens as well as in natural areas. It is a perennial vine that grows quickly, produces copious seeds, and spreads rhizomatously. Researchers at University of Alaska Fairbanks found that compared to five native legume species, bird vetch produced twice the amount of biomass in the presence of both native and non-native soil microbes, suggesting that bird vetch is superior when it comes to nitrogen fixation. Further investigation found that, using only native nitrogen-fixing bacteria, bird vetch produced significantly more root nodules than a native legume species, indicating that it is highly effective at forming relationships with native soil microbes. Additional studies found that the ability of bird vetch to climb up other plants, thereby gaining access to more sunlight and smothering host plants, contributed to its success as an invasive plant.

 Seed pods of bird vetch (Vicia cracca) in Fairbanks, Alaska

Day Three

The final day of the workshop was a veritable cornucopia of topics, including risk assessments for invasive species, profiles of new invasive species, updates on invasive species control projects, discussions about early detection and rapid response (EDRR), and talks about citizen science and community involvement. My head was swimming with impressions and questions. Clearly there are no easy answers when it comes to invasive species, and like other complex, global issues (made more challenging as more players are involved), the increasingly deep well of issues and concerns to resolve is not likely to ever run dry.

Future posts will dig further into some of the discussions that were had on day three. For now, here are a few resources that I gathered throughout the workshop:

Interpretive sign at Alaska Botanical Garden in Anchorage, Alaska

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Seagrass Meadows and Their Role in Healthy Marine Ecosystems

Seagrass meadows are found along soft-bottomed, shallow, marine coastlines of every continent except Antartica. Their abundance and the important roles they play earn them the title of third most valuable ecosystem on the planet after estuaries and wetlands. These extensive meadows are made up of a group of flowering plants that are unique in their ability to thrive submerged in salty seawater. Tossed about by the tides, they feed and harbor an incredibly diverse world of marine life and help protect neighboring ecosystems by stabilizing sediments and mitigating pollution.

Seagrasses are often confused with seaweed, but they are very different organisms. Seaweed is algae. Seagrasses are plants that at one point in their evolutionary history lived on land but then retreated back into the waters of their ancient ancestors. They are rooted in the sediment of the sea floor and, depending on the species, can reproduce both sexually (submerged flowers are pollinated with the help of moving water) and/or asexually (via rhizomes). Although many of them have a grass-like appearance, none of them are in the grass family (Poaceae); instead, the approximately 72 different species belong to one of four families (Posidoniaceae, Zosteraceae, Hydrocharitaceae, or Cymodoceaceae).

Seagrass meadow in Wakaya, Fiji (photo credit: wikimedia commons)

Seagrass meadows can be composed of a single seagrass species or multiple species, with some meadows consisting of a dozen species or more. Seagrasses depend on light for photosynthesis, so they generally occur in shallow areas. How far seagrass meadows extend out into the ocean depends on light availability and the shade tolerance of the seagrass species. Their presence at the shoreline is limited naturally by how exposed they become at low tide, the frequency and strength of waves and associated turbidity, and low salinity from incoming fresh water.

Seagrass meadows benefit life on earth in many ways. As ecosystem engineers they create habitat and produce food for countless species, sequester a remarkable amount of carbon, and help maintain the health of neighboring estuaries, mangroves, coral reefs, and other ecosystems. They are home to commercial fisheries, which provide food for billions of people. Like many ecosystems on the planet, they are threatened by human activity. Pollution, development, recreation, and climate change jeopardize the health and existence of seagrass meadows. Thus, it is imperative that we learn as much as we can about them so that we are better equipped to protect them.

Turtle grass (Thalassia testudinum) growing in an estuary on the coast of San Salvador Island, Bahamas (photo credit: wikimedia commons)

In a report published in a February 2017 issue of Science, researchers examined the ability of seagrass meadows in Indonesia to remove microbial pathogens deposited into the sea via wastewater. When levels of the bacterial pathogen Enterococcus were compared between seagrass meadows and control sites, a three-fold difference was detected, with the seagrass meadows harboring the lowest levels. When other potential disease-causing bacteria were considered, the researchers found that “the relative abundance of bacterial pathogens in seawater” was 50% lower in both the intertidal flat and the coral reefs found within and adjacent to the seagrass meadows compared to control sites.

This has implications for the health of both humans and coral reefs, the latter of which face many threats including bacterial diseases. Two important coral reef diseases, white syndrome and black band disease, as well as signs of mortality associated with bleaching and sediment deposition “were significantly less on reefs adjacent to seagrass meadows compared to paired reefs,” according to the report.

Cushion sea star in seagrass meadow (photo credit: wikimedia commons)

The researchers note that “seagrasses are valued for nutrient cycling, sediment stabilization, reducing the effects of carbon dioxide elevation, and providing nursery habitat for fisheries.” The results of this study demonstrate the potential for seagrass meadows to “significantly reduce bacterial loads,” benefiting “both humans and other organisms in the environment.” Yet another reason to care about and conserve this vital ecosystem.

Additional Resources on Seagrass and Seagrass Conservation:

And if that’s not enough, check out this fun YouTube video:

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When Sunflowers Follow the Sun

Tropisms are widely studied biological phenomena that involve the growth of an organism in response to environmental stimuli. Phototropism is the growth and development of plants in response to light. Heliotropism, a specific form of phototropism, describes growth in response to the sun. Discussions of heliotropism frequently include sunflowers and their ability to “track the sun.” This conjures up images of a field of sunflowers in full bloom following the sun across the sky. However cool this might sound, it simply doesn’t happen. Young sunflowers, before they bloom, track the sun. At maturity and in bloom, the plants hold still.

What is happening in these plants is still pretty cool though, and a report published in an August 2016 issue of Science sheds some light on the heliotropic movements of young sunflowers. They begin the morning facing east. As the sun progresses across the sky, the plants follow, ending the evening facing west. Over night, they reorient themselves to face east again. As they reach maturity, this movement slows, and most of the flowers bloom facing east. Over a series of experiments, researchers were able to determine the cellular and genetic mechanisms involved in this spectacular instance of solar tracking.

Helianthus annuus (common sunflower) is a native of North America, sharing this distinction with dozens of other members of this recognizable genus. It is commonly cultivated for its edible seeds (and the oil produced from them) as well as for its ornamental value. It is a highly variable species and hybridizes readily. Wild populations often cross with cultivated ones, and in many instances the common sunflower is considered a pesky weed. Whether crop, wildflower, or weed, its phototropic movements are easy to detect, making it an excellent subject of study.

Researchers began by tying plants to stakes so that they couldn’t move. Other plants were grown in pots and turned to face west in the morning. The growth of these plants was significantly stunted compared to plants that were not manipulated in these ways, suggesting that solar tracking promotes growth.

The researchers wondered if a circadian system was involved in the movements, and so they took sunflowers that had been growing in pots in a field and placed them indoors beneath a fixed overhead light source. For several days, the plants continued their east to west and back again movements. Over time, the movements became less detectable. This and other experiments led the researchers to conclude that a “circadian clock guides solar tracking in sunflowers.”

Another series of experiments helped the researchers determine what was happening at a cellular level that was causing the eastern side of the stem to grow during the day and the western side to grow during the night. Gene expression and growth hormone levels differed on either side of the stem depending on what time of day it was. In an online article published by University of California Berkeley, Andy Fell summarizes the findings: “[T]here appear to be two growth mechanisms at work in the sunflower stem. The first sets a basic rate of growth for the plant, based on available light. The second, controlled by the circadian clock and influenced by the direction of light, causes the stem to grow more on one side than another, and therefore sway east to west during the day.”

The researchers observed that as the plants reach maturity, they move towards the west less and less. This results in most of the flowers opening in an eastward facing direction. This led them to ask if this behavior offers any sort of ecological advantage. Because flowers are warmer when they are facing the sun, they wondered if they might see an increase in pollinator visits during morning hours on flowers facing east versus those facing west. Indeed, they did: “pollinators visited east-facing heads fivefold more often than west-facing heads.” When west-facing flowers where warmed with a heater in the morning, they received more pollinator visits than west-facing flowers that were not artificially warmed, “albeit [still] fewer than east-facing flowers.” However, increased pollinator visits may be only part of the story, so further investigations are necessary.

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I’m writing a book about weeds, and you can help. For more information, check out my Weeds Poll.

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Love and Hate – The Story of Purple Loosestrife

In the early 1800’s, seeds of purple loosestrife found their way to North America. They arrived from Europe several times by various means – accidentally embedded in the ballast of ships, inadvertently tucked in sheep’s wool, and purposely carried in the hands of humans. Native to much of Europe and parts of Asia and commonly found growing in wetlands and other riparian areas, purple loosestrife’s appealing spikes of magenta flowers, sturdy, upright growth habit, and ease of propagation made it a prized ornamental; its abundant nectar made it a favorite of beekeepers.

During its first 150 years or so in North America, purple loosestrife became naturalized in ditches, wet meadows, and the banks of streams, rivers, lakes, and ponds while also enjoying a place in our gardens. Concern about its spread was raised in the first half of the twentieth century, but it wasn’t until the 1980’s after an extensive survey was done and a special report was issued by the U.S. Fish and Wildlife Service that attitudes about purple loosestrife shifted dramatically. At that point, it was no longer a benign invader and welcome garden companion. It was, instead, a biological menace that needed to be destroyed.

Lytrhrum salicaria – commonly known as purple loosestrife, spiked willow-herb, long purples, rainbow weed, etc. – is an herbaceous perennial in the family Lythraceae. It reaches up to two meters tall; has square or angular stems with lance-shaped, stalkless leaves up to ten centimeters long; and ends in dense, towering spikes of pink-purple, 5-7 petaled flowers. The flowers attract a wide variety of pollinating insects – mostly bees – and afterwards produce small capsules full of tiny, red-brown seeds. Charles Darwin thoroughly studied the flowers of purple loosestrife; he was intrigued by the plant for many reasons, including its heterostyly (a topic for another post).

Lythrum salicaria (purple loosestrife) – image credit: wikimedia commons

Purple loosestrife seeds remain viable for up to 20 years and are transported by wind, water, and in mud stuck to the feet of birds. Apart from seeds, populations expand clonally as root crowns grow larger each year and produce increasingly more stems. Broken stem pieces also take root in mud, creating new plants. Purple loosestrife’s ability to form expansive populations in a quick manner, pushing other plants aside and forming what appears to be a dense monoculture, is part of the reason it has earned itself a place among the International Union for Conservation of Nature’s list of 100 World’s Worst Invasive Alien Species.

But is this ranking justified? In a paper published in Biological Invasions in 2010, Claude Lavoie compares news reports about purple loosestrife around the turn of the century with data presented in scientific papers and finds that the reports largely exaggerate the evidence. Purple loosestrife was being accused of all manner of crimes against nature and was being condemned before there was sound evidence to justify such actions.

It began with the U.S. Fish and Wildlife Service’s special report published in 1987. According to Lavoie, “a long list of the impacts of the species on wetland flora and fauna [was] presented,” but the claims were not supported by observational or experimental data – “the impacts [were] only suspected.” Regardless, wetland managers began campaigns against purple loosestrife in order to convince the public that it was a Beautiful Killer. News outlets were quick to spread the word about this “killer” plant. When biological control programs began in the 1990’s, news outlets reported on their success. Little empirical evidence had been published on either topic, and debates about purple loosestrife’s impacts remained unsettled in the scientific community.

The flowers of purple loosestrife (Lythrum salicaria) – photo credit: wikimedia commons

Around this time, five reviews were published examining the evidence against purple loosestrife. Lavoie reports that all but one of them “rely on a relatively high number of sources that have not been published in peer-reviewed journals.” After examining the reviews, Lavoie concludes: “although each review provided valuable information on purple loosestrife, most were somewhat biased and relied on a substantial amount of information that was anecdotal or not screened by reviewers during a formal evaluation process. Only one review was impartial, and this one painted an inconclusive picture of the species.”

Research has continued regarding the impacts of purple loosestrife, and so Lavoie examined 34 studies that were published during the 2000’s in search of conclusive evidence that the plant is as destructive to wetlands and wildlife as has been claimed. Upon examination he concludes that “stating that this plant has ‘large negative impacts’ on wetlands is probably exaggerated.” The most common accusation – that purple loosestrife crowds out native plants and forms a monoculture – “is controversial and has not been observed in nature (with maybe one exception).” Lavoie finds that there is “certainly no evidence that purple loosestrife ‘kills wetlands’ or ‘creates biological deserts,'” and “there are no published studies [in peer-reviewed journals] demonstrating that purple loosestrife has an impact on waterfowl or fishes.” All other negative claims against purple loosestrife “have not been the object of a study,” except for its impact on amphibians, which had at that time only been tested on two species, one “reacting negatively.” Certain claims – such as purple loosestrife’s impact on wetland hydrology – should be studied more in depth “considering the apparent public consensus on the detrimental effects of purple loosestrife” on wetland ecosystems.

Lavoie agrees that it is reasonable to control purple loosestrife when the intention is to reduce additional pressures on an ecosystem that is already highly threatened. However, he warns that “focusing on purple loosestrife instead of on other invasive species or on wetland losses to agriculture or urban sprawl could divert the attention of environmental managers from more urgent protection needs.” There is mounting evidence that purple loosestrife invasions are disturbance-dependent and are “an indicator of anthropogenic disturbances.” In order to protect our wetlands, we must first protect them against undue disturbance. Lavoie supports using the Precautionary Principle when dealing with introduced species; however, he finds the approach “much more valuable for newcomers than for invaders coexisting with native species for more than a century.”

A field of purple loosestrife in Massachusetts – photo credit: wikimedia commons

Purple loosestrife has found its way to nearly every state in America and most of the Canadian provinces. Peter Del Tredici writes in Wild Urban Plants of the Northeast, “Conservationists despise purple loosestrife, despite its beauty, and it is listed as an invasive species in most of the states where it grows.” By listing a plant as a noxious weed, landowners are obligated to remove it. Care must be taken though, as removal of purple loosestrife can result in a secondary invasion by noxious weeds with an even worse track record, such as common reed or reed canary grass. “Hardly a gain from the biodiversity point of view,” quips Lavoie.

Claude Lavoie’s paper and the papers he references are definitely worth reading. It is important that we continue to study purple loosestrife and species like it in order to fully understand the impact that introduced species are having on natural areas, especially since it is unlikely that we will ever completely eliminate them. On that note, I’ll leave you with this passage from The Book of Swamp and Bog by John Eastman:

The situation is easy for environmentalists to deplore. This plant, like few others, stirs our alien prejudice. Our native cattails, for example, are almost as rudely aggressive and competitive in many wetland areas as purple loosestrife. Yet, because cattails obvioulsy ‘belong here,’ they seldom evoke the same outraged feelings against their existence. … With the spread of purple loosestrife, we have new opportunities to witness the phases of an ever-recurring ecological process. We can watch it affect, change, adapt, and refit both its own elements and those of invaded communities into new arrangements of energy efficiency. The point is that we might as well study this process rather than simply deplore it; we have few alternatives.

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Screening for Invasive Plants at Botanical Gardens and Arboreta

As discussed in last week’s post, many of the invasive species that we find in our natural areas were first introduced to North America via the horticulture trade. As awareness of this phenomenon grows, steps are being taken by the horticulture industry to address this issue. The concluding remarks by Sarah Reichard and Peter White in their 2001 article in BioScience describe some recommended actions. One of them involves the leadership role that botanical gardens can play by both stopping the introduction and spread of invasive species and by presenting or promoting public education programs.

Reichard and White offer North Carolina Botanical Garden as an example, citing their “Chapel Hill Challenge,” which urges botanical gardens to “do no harm to plant diversity and natural areas.” Reichard and White also encourage botanical gardens and nurseries to adopt a code of conservation ethics addressing invasive species and other conservation issues. Codes of conduct for invasive species have since been developed for the botanical garden community and are endorsed by the American Public Gardens Association.

 

Botanical gardens that adopt this code have a number of responsibilities, one of which is to “establish an invasive plant assessment procedure,” preferably one that predicts the risks of plant species that are new to the gardens. In other words, botanical gardens are encouraged to screen the plants that are currently in their collections, as well as plants that are being added, to determine whether these plants currently exhibit invasive behavior or have the potential to become invasive. Many botanical gardens now have such programs in place, and while they may not be able to predict all invasions, they are a step in the right direction.

In an article published in Weed Technology (2004), staff members at Chicago Botanic Garden (CBG) describe the process they went through to determine a screening process that would work for them. CBG has an active plant exploration program, collecting plants in Asia, Europe, and other parts of North America. Apart from adding plants to their collection, one of the goals of this program is to find plants with horticulture potential and, through their Ornamental Plant Development department, prepare these plants to be introduced to the nursery industry in the Chicago region. As their concern about invasive species has grown, CBG (guided by a robust Invasive Plant Policy) has expanded and strengthened its screening process.

In order to do this, CBG first evaluated three common weed risk assessment models. The models were modified slightly in order to adapt them to the Chicago region. Forty exotic species (20 known invasives and 20 known non-invasives) were selected for testing. Each invasive was matched with a noninvasive from the same genus, family, or growth form in order to “minimize ‘noise’ associated with phylogenetic differences.” The selected species also included an even distribution of forbs, vines, shrubs, and trees.

Weed risk assessment models are used to quickly determine the potential of a plant species to become invasive by asking a series of questions about the plant’s attributes and life history traits, as well as its native climate and geography. A plant species can be accepted, rejected, or require further evaluation depending on how the questions are answered. For example, if a plant is known to be invasive elsewhere and/or if it displays traits commonly found in other invasive species, it receives a high score and is either rejected or evaluated further. Such models offer a quick and affordable way to weed out incoming invasives; however, they are not likely to spot every potential invasive species, and they may also lead to the rejection of species that ultimately would not have become invasive.

After testing the three models, CBG settled on the IOWA-modified Reichard and Hamilton model “because it was extensively tested in a climatic zone reasonably analogous to … Illinois,” and because it is easy to use and limits the possibility of a plant being falsely accepted or rejected. The selected model was then tested on 208 plants that were collected in the Republic of Georgia. Because few details were known about some of the plants, many of the questions posed by the model could not be answered. This lead CBG to modify their model to allow for such plants to be grown out in quarantined garden plots. This way pertinent information can be gathered, such as “duration to maturity; self-compatibility; fruit type and potential methods of seed or fruit dispersal; seed production, viability, and longevity in the field; and vegetative spread.” CBG believes that evaluations such as this will help them modify their model over time and give them more confidence in their screening efforts.

More about botanical gardens and invasive species: Botanic Gardens Conservation International – Invasive Alien Species

More about weed risk assessment models: Weed Risk Assessment – A way forward or a waste of time? by Philip E. Hulme

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When Alien Plants Invade – The Four Stages of Invasion, part one

As humans move around the globe, they are regularly accompanied by plants. Some plant species are intentional guests, while others are interlopers. This steady movement of plants from one region to another results in plants being introduced to areas where they are not native. In this regard, they are aliens. Some of these alien species will take up permanent residence and, as a result, can disrupt ecosystems, compete with native plant species, and cause economic damage. This earns them the title “invasive”. But not all introduced plant species achieve this. In fact, many will find themselves in a new region but will be unable to colonize. Others will colonize but not become fully established. Still others become established but will not spread. In all cases there are factors at play that either aid or limit an introduced plant species in becoming invasive.

In a review published in New Phytologist (2007), Kathleen Theoharides and Jeffrey Dukes examine four stages of invasion (transport, colonization, establishment, and landscape spread) and some of the “filters” that occur in each stage that help determine whether or not an alien plant species will become invasive. In their introduction they clarify, “these stages are not discrete, and filters will likely affect more than one stage,” but by analyzing each of the stages we can better determine how and why some introduced species are successful at becoming invasive while others are not. Generalities derived from this investigation can “be used to predict the outcome of invasion events, or to explore mechanisms responsible for deviations from these generalizations.”

Theoharides, K. A. and Dukes, J. S. (2007), Plant invasion across space and time: factors affecting nonindigenous species success during four stages of invasion. New Phytologist, 176: 256–273. doi:10.1111/j.1469-8137.2007.02207.x

In part one, we will look at the first two stages of invasion: transport and colonization.

Species have always moved around from region to region by various means. However, as Theoharides and Dukes write, “current species movements are happening faster than before and from more distant regions, primarily as a result of global commerce and travel.” When it comes to human-mediated dispersal, many plants may never be transported by humans, while others simply won’t survive the journey. Species that are widespread may have a better chance of being transported because they are more likely to make contact with humans. Transporting high numbers of propagules (i.e. seeds, spores, cuttings) generally increases the likelihood that a species will survive the journey.

The invasion of Phalaris arundinacea (reed canary grass) was facilitated by multiple introduction events from a variety of sources within the native European range.” — Theoharides, K. A. and Dukes, J. S. (2007) [photo credit: eol.org]

Plants are transported by humans for many reasons. Sometimes its accidental, but often it is purposeful for either utilitarian or aesthetic purposes. Plants provide us with food, fuel, forage, building materials, clothing, and medicine. Over millennia, we have selected suites of species that are ideal for such purposes, and we have carried them with us into new regions or brought them home from other parts of the world. Not all of these species are well-behaved in their new homes, and many have become invasive. These species are given an advantage because they have been selected for traits like cold hardiness, disease resistance, and high yield. When they are transported, they are brought to locations with similar climates. “Climate matching, combined with intentional cultivation, greatly increases the likelihood that [these] species will escape cultivation.”

Surviving transportation is not a guarantee that alien plants will successfully colonize a new area. Myriad environmental conditions and biological processes stand in their way. Much depends on propagule pressure – “the combined measure of the number of individuals reaching a new area in any one release event and the number of discrete release events.” Where propagule pressure is high, colonization is more likely. Repeated introductions across a large area offer the species a greater chance of finding itself in a suitable location as well as a greater level of genetic variation. Disturbed environments with less competitors and increased resources (i.e. light, moisture, soil nutrients) are often easier to colonize than locations with a high level of biodiversity and fewer available resources.

“Populations of Salix babylonica (weeping willow) in New Zealand may have invaded from a single cutting.” — Theoharides, K. A. and Dukes, J. S. (2007) [photo credit: eol.org]

Climate is one of the main filters of colonization, yet plant species have still managed to colonize regions with very different climates compared to what they’re used to, while other plant species have been unsuccessful in colonizing regions with similar climates. Plant species that originate from wide geographic ranges tend to have “broader climatic tolerances” – a trait that along with phenotypic plasticity and a high level of genetic variability can enable a species to adapt to new and challenging environments. Other advantageous traits include “fast growth, self-compatibility, a short juvenile period, and seeds that germinate without a pre-treatment.”

If and when colonization is achieved, establishment is no guarantee. “In order for a plant to establish itself it must continue to increase from low density over the long term.” Small numbers of plants may successfully reproduce, but environmental factors, genetic issues, and biological competition may still stand in their way. Species that invade disturbed sites where resource availability is temporarily high, may soon find themselves in a resource-limited situation. As a result, their populations may dwindle.

With transport and colonization accomplished, establishment is the next goal. Establishment and landscape spread will be covered in part two.

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Seed Dispersal via Caching – The Story of Antelope Bitterbrush

Generally speaking, individual plants produce an enormous amount of seeds. This may seem like a huge waste of resources, but the reality is that while each seed has the potential to grow into an adult plant that will one day produce seeds of its own, relatively few may achieve this. Some seeds will be eaten before they get a chance to germinate. Others germinate and soon die from lack of water, disease, or herbivory. Those that make it past the seedling stage continue to face similar pressures. Reaching adulthood, then, is a remarkable achievement.

Antelope bitterbrush is a shrub that produces hundreds of seeds per individual. Each seed is about the size of an apple seed. Some seeds may be eaten right away. Others fall to the ground and are ignored. But a large number are collected by rodents and either stored in burrows (larder hoarding) or in shallow depressions in the soil (scatter hoarding). It is through caching that antelope bitterbrush seeds are best dispersed. When rodents fail to return to caches during the winter, the seeds are free to sprout in the spring. Some of the seedlings will dry out and others will be eaten, but a few will survive, making the effort to produce all those seeds worth it in the end.

Fruits forming on antelope bitterbrush (Purshia tridentata)

Antelope bitterbrush (Purshia tridentata) is in the rose family and is often simply referred to as bitterbrush. It occurs in grasslands, shrub steppes, and dry woodlands throughout large sections of western North America. It is a deciduous shrub that generally reaches between three and nine feet tall but can grow up to twelve feet. It has wedge-shaped leaves that are green on top, grayish on bottom, and three-lobed. Flowers are yellow, strongly fragrant, and similar in appearance to others in the rose family. Flowering occurs mid-spring to early summer. Fruits are achenes – single seeds surrounded by papery or leathery coverings. The covering must rot away or be removed by animals before the seed can germinate.

Bitterbrush is an important species for wildlife. It is browsed by mule deer, pronghorn antelope, bighorn sheep, and other ungulates, including livestock. It provides cover for birds, rodents, reptiles, and ungulates. Its seeds are collected by harvester ants and rodents, its foliage is consumed by tent caterpillars and other insects, and its flowers are visited by a suite of pollinators. For all that it offers to the animal kingdom, it also relies on it for pollination and seed dispersal. The flowers of bitterbrush are self-incompatible, and if it wasn’t for ants and rodents, the heavy seeds – left to rely on wind and gravity – would have trouble getting any further than just a few feet from the parent plant.

Antelope bitterbrush (Purshia tridentata) in full bloom – photo credit: wikimedia commons

In a study published in The American Naturalist (February 1993), Stephen Vander Wall reported that yellow pine chipmunks were the primary dispersal agents of bitterbrush seeds in his Sierra Nevada study area. The optimal depth for seedling establishment was between 10-30 millimeters. Seeds that are cached too near the surface risk being pushed out of the ground during freeze and thaw cycles where they can desiccate upon germination. Cached bitterbrush seeds benefit when there are several seeds per cache because, as Vander Wall notes, “clumps of seedlings are better able to push through the soil and can establish from greater depths than single seedlings.”

Another study by Vander Wall, published in Ecology (October 1994), reiterated the importance of seed caching by yellow pine chipmunks in the establishment of bitterbrush seedlings. Seed caches, which consisted of anywhere from two to over a hundred seeds, were located as far as 25 meters from the parent plant. Cached seeds are occasionally moved to another location, but Vander Wall found that even these secondary caches produce seedlings. Of course, not all of the seedlings that sprout grow to maturity. Vander Wall states, “attrition over the years gradually reduces the number of seedlings within clumps.” Yet, more than half of the mature shrubs he observed in his study consisted of two or more individuals, leading him to conclude that “they arose from rodent caches.”

A study published in the Journal of Range Management (January 1996) looked at the herbivory of bitterbrush seedlings by rodents. In the introduction the authors discuss how “rodents [may] not only benefit from antelope bitterbrush seed caches as a future seed source, but also benefit from the sprouting of their caches as they return to graze the cotyledons of germinating seeds.”  In this study, Ord’s kangaroo rats, deer mice, and Great Basin pocket mice were all observed consuming bitterbrush seedlings, preferring them even when millet was offered as an alternative. The two species of mice also dug up seedlings, possibly searching for ungerminated seeds. Despite seed dispersal via caching, an overabundance of rodents can result in few bitterbrush seedlings reaching maturity.

A cluster of antelope bitterbrush seedlings that has been browsed. “Succulent, young seedlings are thought to be important in the diets of rodents during early spring because of the nutrients and water they contain.” — Vander Wall (1994)

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Photos of antelope bitterbrush seedling clusters were taken at Idaho Botanical Garden, where numerous clusters are presently on display along the pathways of the native plant gardens and the adjoining natural areas. 

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Alien Plant Invasions and the Extinction Trajectory

One of the concerns about introduced species becoming invasive is that they threaten to reduce the biodiversity of the ecosystems they have invaded. They do this by spreading rampantly, using up resources and space, altering ecosystem functions, and ultimately pushing other species out. In the case of certain invasive animals, species may be eliminated via predation; but plants don’t eat each other (generally), so if one plant species is to snuff out another plant species it must use other means. Presently, we have no evidence that a native plant species has been rendered extinct solely as a result of an invasive plant species. That does not mean, however, that invasive plants are not doing harm.

In a paper published in AoB Plants in August 2016, Paul O. Downey and David M. Richardson argue that, when it comes to plants, focusing our attention on extinctions masks the real impact that invasive species can have. In general, plants go extinct more slowly than animals, and it is difficult to determine that a plant species has truly gone extinct. Some plants are very long-lived, so the march towards extinction can extend across centuries. But the real challenge – after determining that there are no above-ground signs of life – is determining that no viable seeds remain in the soil (i.e. seed bank). Depending on the species, seeds can remain viable for dozens (even hundreds) of years, so when conditions are right, a species thought to be extinct can emerge once again. (Consider the story of the Kankakee mallow.)

On the other hand, there is plenty of evidence that invasive plant species have had significant impacts on certain native plant populations and have placed such species on, what Downey and Richardson call, an extinction trajectory. It is this trajectory that deserves our attention if our goal is to save native plant species from extinction. As described in the paper, the extinction trajectory has six steps – or thresholds – which are defined in the infographic below:

6-threshold-extinction-trajectory

Downey and Richardson spend a portion of the paper summarizing research that demonstrates how invasive plants have driven native plants into thresholds 1-3, thereby placing them on an extinction trajectory. In New Zealand, Lantana camara (introduced from the American tropics) creates dense thickets, outcompeting native plants. Researchers found that species richness of native plants declined once L. camara achieved 75% cover in the test sites. In the U.S., researchers found reduced seed set in three native perennial herbs as a result of sharing space with Lonicera mackii (introduced from Asia), suggesting that the alien species is likely to have a negative impact on the long-term survivability of these native plants. Citing such research, Downy and Richardson conclude that “it is the direction of change that is fundamentally important – the extinction trajectory and the thresholds that have been breached – not whether a native plant species has actually been documented as going extinct due to an alien plant species based on a snapshot view.”

Introduced to New Zealand from the American tropics, largeleaf lantana (Lantana camara) forms dense thickets that can outcompete native plant species. (photo credit: wikimedia commons)

Introduced to New Zealand from the American tropics, largeleaf lantana (Lantana camara) forms dense thickets that can outcompete native plant species. (photo credit: wikimedia commons)

In support of their argument, they also address problems with the way some research is done (“in many instances appropriate data are not collected over sufficiently long periods,” etc.), and they highlight the dearth of data and research (“impacts associated with most invasive alien plants have not been studied or are poorly understood or documented”). With those things in mind, they make recommendations for improving research and they encourage long-term studies and collaboration in order to address the current “lack of meta analyses or global datasets.” A similar recommendation was made in American Journal of Botany in June 2015.

The language in this report makes it clear that the authors are responding to a certain group of people that have questioned whether or not the threat of invasive plants has been overstated and if the measures we are taking to control invasive plants are justified. The following cartoon that appeared along with a summary of the article way oversimplifies the debate:

04_figure2

Boy: There are no studies that show weeds cause native plants to go extinct, thus we should not control them. Plant: If we wait until then, we’ll all be gone!!! Girl: Just because no one has demonstrated it does not mean that extinctions do not occur. The problem is not overstated!

It seems to me that a big part of why we have not linked an invasive plant species to a native plant species extinction (apart from the difficulty of determining with certainty that a plant has gone extinct) is that extinctions are often the result of a number of factors. The authors do eventually say that: “it is rare that one threatening process in isolation leads to the extinction of a species.” So, as much as it is important to fully understand the impacts that invasive plant species are having, it is also important to look at the larger picture. What else is going on that may be contributing to population declines?

Observing invaded plant populations over a long period seems like our best bet in determining the real effects that invasive species are having. In some cases, as Downey and Richardson admit, “decreased effects over time” have been documented, and so “the effects [of invasive species] are dynamic, not static.” And speaking of things that are dynamic, extinction is a dynamic process and one that we generally consider to be wholly negative. But why? What if that isn’t always the case? Extinctions have been a part of life on earth as long as life has been around. Is there anything “good” that can come out of them?